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    275RESONANCE March 2013

    GENERAL ARTICLE

    Plant Growth Promoting Rhizobacteria

    Potential Microbes for Sustainable Agriculture

    Jay Shankar Singh

    Keywords

    Agriculture, biofertilizers, PGPR,

    sustainable agriculture.

    Plant Growth Promoting Rhizobacteria (PGPR) are a group

    of bacteria that enhances plant growth and yield via various

    plant growth promoting substances as well as biofertilizers.

    Given the negative environmental impact of artificial fertiliz-

    ers and their increasing costs, the use of beneficial soil micro-

    organisms such as PGPR for sustainable and safe agriculture

    has increased globally during the last couple of decades.

    PGPR as biofertilizers are well recognized as efficient soil

    microbes for sustainable agriculture and hold great promise

    in the improvement of agriculture yields.

    Agriculture contributes to a major share of national income and

    export earnings in many developing countries, while ensuring

    food security and employment. Sustainable agriculture is vitally

    important in todays world because it offers the potential to meet

    our future agricultural needs, something that conventional agri-

    culture will not be able to do. Recently there has been a great

    interest in eco-friendly and sustainable agriculture. PGPR areknown to improve plant growth in many ways when compared to

    synthetic fertilizers, insecticides and pesticides. They enhance

    crop growth and can help in sustainability of safe environment

    and crop productivity. The rhizospheric soil1 contains diverse

    types of PGPR communities, which exhibit beneficial effects on

    crop productivity. Several research investigations are conducted

    on the understanding of the diversity, dynamics and importance

    of soil PGPR communities and their beneficial and cooperative

    roles in agricultural productivity. Some common examples of

    PGPR genera exhibiting plant growth promoting activity are:

    Pseudomonas, Azospirillum, Azotobacter, Bacillus, Burkholdaria,

    Enterobacter, Rhizobium, Erwinia, Mycobacterium, Mesorhizo-

    bium, Flavobacterium, etc. This article presents perspectives on

    the role of PGPR in agriculture sustainability.

    Jay Shankar Singh is an

    Assistant Professor in the

    Department of Environ-

    mental Microbiology, BB

    Ambedkar (Cen tral)

    University, Luc know,

    Uttar Pradesh, India. He is

    orking on the environ-

    mental problems related to

    ecology of methane (CH4)-

    consuming bacteria

    (methanotrophs),

    environmental remediation

    and sustainable agricul-

    ture development. He also

    has a strong devotion to

    teaching and is always

    available to share with

    others the ever-growing

    enthusiasm for his

    research field.

    1 The soil surrounding plant root

    area where intense biological

    and chemical activities are influ-

    enced by root exudates, and mi-

    croorganisms f eeding on the

    compounds.

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    GENERAL ARTICLE

    PGPR as Biofertilizers

    Free-living PGPR have shown promise as biofertilizers. Many

    studies and reviews have reported plant growth promotion, in-

    creased yield, solubilization of P (phosphorus) or K (potassium),

    uptake of N (nitrogen) and some other elements through inocula-

    tion with PGPR. In addition, studies have shown that inoculation

    with PGPR enhances root growth, leading to a root system with

    large surface area and increased number of root hairs.

    A huge amount of artificial fertilizes are used to replenish soil N

    and P, resulting in high costs and increased environmental pollu-

    tion. Most of P in insoluble compounds is unavailable to plants.

    N2-fixing and P-solubilizing bacteria may be important for plantnutrition by increasing N and P uptake by the crop plants, and

    playing a crucial role in biofertilization. N2-fixation and P-solubi-

    lization, production of antibiotics, and other plant growth pro-

    moting substances are the principal contribution of the PGPR in

    the agro-ecosystems. More recent research findings indicate that

    the treatment of agricultural soils with PGPR inoculation signifi-

    cantly increases agronomic yields as compared to uninoculated

    soils.

    Mechanisms of Plant Growth Promotion by PGPR

    Several mechanisms have been suggested by which PGPR can

    promote plant growth; some important ones are as follows (Table 1):

    Box 1.

    Plant growth promoting rhizobacteria (PGPR): PGPR (rhizo-biofertilizers) are a group of bacteria that

    actively colonize plant roots and enhance plant growth and yield.

    Biofertilizer: Composition of living micro-organisms which, when applied to surface of plant, seed, or soil,

    colonize the plant rhizosphere or inside the plant body (as endophyte) and promotes plant growth through

    enhancing the supply or availability of fundamental nutrients to crop plants.

    Sustainable agriculture: Sustainable agriculture involves the successful management of agricultural re-

    sources to satisfy human needs while maintaining or enhancing environmental quality without exploiting the

    natural resources of future generations.

    Many studies and

    reviewshave reported

    plant growthpromotion, increased

    yield, solubilization of

    P (phosphorus) or K

    (potassium), uptake

    of N (nitrogen), etc.

    through inoculation

    with PGPR.

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    Phytohormone Production

    The enhancement in various agronomic yields due to PGPR has

    been reported because of the production of growth stimulating

    phytohormones (Table 2) such as indole-3-acetic acid (IAA),

    gibberellic acid (GA3), zeatin, ethylene and abscisic acid (ABA).

    PGPR Crop parameters

    Rhizobium leguminosarum Direct growth promotion of canola and lettuce

    Pseudomonas putida Early developments of canola seedlings, growth

    stimulation of tomato plant

    Azospirillum brasilense andA. irakense Growth of wheat and maize plants

    P. flurescens Growth of pearl millet, increase in growth, leaf

    nutrient contents and yield ofbanana (Musa)

    Azotobacterand Azospirillum spp. Growth and productivity of canola

    P. alcaligenes, Bacillus polymyxa, Enhances uptakeof N, P and K bymaize crop

    andMycobacterium phlei

    Pseudomonas, Azotobacte rand Stumulates growth and yield of chick pea (Cicer

    Azospirillum spp. arietinum)

    R. leguminismarum and Improves the yield and phosphor us upt ake in wheat

    Pseudomonas spp.

    P. putida, P. flurescens, A. brasilense Improves seed germination, seedling growth and

    andA. lipoferum yield of maize

    P. putida, P. fluoresce ns, P. fluorescens, Improves seed germination, growth parameters of

    P. putida, A. lipoferum, A. brasilense maize seedling in greenhouse and also grain yield

    of field grown maize

    Table 1. PGPR and their ef-

    fect on growth parameters/

    yields of crop/fruit plants.

    Phytohormones PGPR

    Indole-3-acetic acid (IAA) Acetobacter diazotrophicus

    andHerbaspirillum seropedicae

    Zeatin and ethylene Azospirillum sp.

    Gibberellic acid (GA3) Azospirillum lipoferum

    Abscisic acid (ABA) Azospirillum brasilense

    Table 2. Examples of differ-

    ent phytohormone-producing

    PGPR.

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    Recent studies confirm that the treatment of seeds or cuttings with

    non-pathogenic bacteria, such as Agrobacterium, Bacillus, Strep-

    tomyces, Pseudomonas, Alcaligenes, etc. induce root formationin some plants because of natural plant growth promoting sub-

    stances produced by the bacteria. Although the mechanisms are

    not completely understood, root induction by PGPR is the ac-

    cepted result of phytohormones such as auxins, growth inhibiting

    ethylene and mineralization. Environment-friendly applications

    in agriculture have gained more importance, in particular in

    horticulture and nursery production. The use of PGPR for nursery

    material multiplication may be important for obtaining organic

    nursery material because the use of all formulations of synthetic

    plant growth regulators, such as indole-3-butyric acid (IBA), are

    prohibited in organic agriculture throughout the world.

    Phosphate Solubilization

    Rhizobium and phosphorus (P) solubilizing bacteria are impor-

    tant to plant nutrition. These microbes also play a significant role

    as PGPR in the biofertilization of crops. These bacteria secrete

    different types of organic acids (e.g., carboxylic acid) thus lower-

    ing the pH in the rhizosphere and consequently release the bound

    forms of phosphate like Ca3

    (PO4)

    2in the calcareous soils. Utili-

    zation of these microorganisms as environment-friendlybiofertilizer helps to reduce the use of expensive phosphatic

    fertilizers. Phosphorus biofertilizers could help increase the avail-

    ability of accumulated phosphate (by solubilization), increase the

    efficiency of biological nitrogen fixation and render availability

    of Fe, Zn, etc., through production of plant growth promoting

    substances.

    Siderophore Production

    PGPR are reported to secrete some extracellular metabolites

    called siderophores. For the first time Kloepper et al. (1980)reported the significance of siderophores produced by certain

    genera of PGPR in plant growth promotion. Siderophores are

    commonly referred to as microbial Fe-chelating low molecular

    The enhancement in

    agronomic yields by

    PGPR is due toproduction of growth

    stimulating

    phytohormones and

    the consequent

    release of the bound

    form of phosphate,

    andextracellular

    metabolites.

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    GENERAL ARTICLE

    weight compounds. The presence of siderophore-producing PGPR

    in rhizosphere increases the rate of Fe3+ supply to plants and

    therefore enhance the plant growth and productivity of crop.Further, this compound after chelating Fe3+ makes the soil Fe3+

    deficient for other soil microbes and consequently inhibits the

    activity of competitive microbes.

    PGPR as Biocontrol Agents

    PGPR produce substances that also protect them against various

    diseases. PGPR may protect plants against pathogens by direct

    antagonistic interactions between the biocontrol agent and the

    pathogen, as well as by induction of host resistance. In recent

    years, the role of siderophore-producing PGPR in biocontrol ofsoil-borne plant pathogens has created great interest. Microbiolo-

    gists have developed techniques for introduction of siderophore-

    producing PGPR in soil system through seed, soil or r oot system.

    PGPR that indirectly enhance plant growth via suppression of

    phytopathogens do so by a variety of mechanisms. These include:

    The ability to produce siderophores (as discussed above) thatchelate iron, making it unavailable to pathogens.

    The capacity to synthesize anti-fungal metabolites such as

    antibiotics, fungal cell wall-lysing enzymes, or hydrogen cya-nide, which suppress the growth of fungal pathogens.

    The ability to successfully compete with pathogens for nutri-ents or specific niches on the root; and the ability to induce

    systemic resistance.

    Among the various PGPRs identified, Pseudomonas fluorescens

    is one of the most extensively studied rhizobacteria, because of its

    antagonistic action against several plantpathogens. Banana bunchy

    top virus (BBTV) is one of the deadly virus which severely

    affects the yield of banana (Musa spp.) crop in Western Ghats,Tamil Nadu, India. It has been demonstrated that application of

    P. fluorescens strain significantly reduced the BBTV incidence

    in hill banana under greenhouse and field conditions. Different

    PGPR produce

    substances that

    also protect themagainst various

    diseases

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    GENERAL ARTICLE

    Table3. PGPR as biocontrol

    agents against various plant

    diseases.

    PGPR Disease resistance

    Bacillus pumilus, Kluyvera cryocrescens, Cucumber Mosaic Cucumovirus (CMV) of tomato

    B. amyloliquef aciens and B. subtilus (Lycopersicon esculentum)

    B. amylolique faciens, B. subtilis and Tomato mottle virus

    B. pumilus

    B. pumilus Bacterial wilt disease in cucumber (Cucumis

    sativus), Blue mold disease of tobacco ( icotiana)

    Pseudomonas fluorescens Sheath blight disease and leaf folder insect in rice

    (Oryza sativa), Reduce the Banana Bunchy Top

    Virus (BBTV) incidence, Saline resistance in

    groundnut (Arachis hypogea)

    B . subtilis and B. pumilus Downy mildew in pearl millet (Pennisetum

    glaucum)

    B. subtilis CMVin cucumber

    B. cereus Foliar diseases of tomato

    Bacillus spp. Blight of bell pepper (Capsicum annuum), Blight

    of squash

    Burkholderia Maize (Zea mays)rot

    B. subtilis Soil borne pathogen of cucumber and pepper (Piper)

    Bacillus sp. andAzospirillum Rice blast

    Fluorescent Pseudomonas spp. Rice sheath rot (Sarocladium oryzae)

    PGPR species as biocontrol agents against various plant diseases

    are given in Table 3.

    PGPR as Biological Fungicides

    PGPR and bacterial endophytes play a vital role in the manage-

    ment of various fungal diseases (Box 2). But one of the major

    hurdles experienced with biocontrol agents is lack of appropriate

    delivery system.

    PGPR and bacterial

    endophytes play a

    vital role in themanagement of

    various fungal

    diseases.

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    GENERAL ARTICLE

    Suggested Reading

    [1] B R Glick, C L Patten, G Holguin and D M Penrose, Biochemical and genetic

    mechanisms used by plant growth promoting bacteria, Imperial College Press,

    London. 1999.

    [2] A R Podile and K Kishore, Plant growth-promoting rhizobacteria. In: Plant

    associated bacteria. Springer Netherlands, pp. 195230, 2007.

    Conclusion

    Worldwide, considerable progress has been achieved in the area

    of PGPR biofertilizer technology. It has been also demonstratedand proved that PGPR can be very effective and are potential

    microbes for enriching the soil fertility and enhancing the agricul-

    ture yield. PGPR are excellent model systems which can provide

    the biotechnologist with novel genetic constituents and bioactive

    chemicals having diverse uses in agriculture and environmental

    sustainability. Current and future progress in our understanding of

    PGPR diversity, colonization ability, mechanisms of action, for-

    mulation, and application could facilitate their development as

    reliable components in the management of sustainable agricul-

    tural systems.

    Address for Correspondence

    Jay Shankar Singh

    Department of Environment

    Microbiology

    School for Environmental

    Sciences, BB Ambedkar

    (Central) University,

    Raibarely R oad, Lucknow

    226025

    Uttar Pradesh, India.

    Email:

    jayshank ar_1@ yahoo.co.i n

    Box 2

    Bacillu s subt ilis , Pseudomonas chlororaphis, endophytic P. fluo rescens inhibit the growth of stem blightpatho gen Corynespora casiicola. The seed treatment and soil application of P. fluorescens reduce root rot

    of black gram caused by Macrophomina phaseolina. Seed and foliar application of P. fluorescens reduce

    sheath blight of rice. B. subtilis in peat supplemented with chitin or chitin-containing materials show better

    control of Aspergillus niger and Fusarium udum in groundnut and pigeon pea, respectively. Strains of

    Burkholder ia cepa cia have been shown to have biocontrol of Fusarium spp.

    It has been also

    demonstrated and

    proved that PGPR

    can be very

    effective and are

    potential microbes

    for enriching the soil

    and enhancing the

    agricultureyield.